Showing papers in "Systematic Entomology in 2017"

The peril of dating beetles

Abstract: Recently, McKenna et al., 2015 (MCK15 hereafter) investigated the higher level phylogenetic relationships of beetles (Insecta, Coleoptera) using the most comprehensive molecular dataset to date, and inferred the absolute ages of major groups using multiple fossil calibrations across the beetle tree of life. Based on the result of their dating analysis, beetles diverged from Strepsiptera in the Early Permian c. 278.33 Ma with a 95% credibility interval (95% CI) of 288.28 to 271.89 Ma, and the crown age of Coleoptera was estimated for the Late Permian c. 252.89 Ma (95% CI: 267.68 to 238.78 Ma), supporting the view that beetles originated before and survived through the End-Permian Mass Extinction that occurred c. 252 Ma (Shen et al., 2011). However, some of the age estimates found in MCK15 are in conflict with current knowledge of the beetle fossil record (e.g. Nikolajev & Ren, 2010; Pan et al., 2011, Prokin & Ren, 2011; Fikáček et al., 2012a; Wang et al., 2013, 2014; Cai et al., 2014b, 2015a; Kirejtshuk et al., 2014; Boucher et al., 2016) and with other recently published molecular age estimates for some major beetle clades (e.g. Zhang & Zhou, 2013; Ahrens et al., 2014; Bloom et al., 2014; Kergoat et al., 2014; Kim & Farrell, 2015; Bocák et al., 2016; Gunter et al., 2016). In some cases, the difference in age estimates is significant and might change our understanding of the mode and tempo of diversification dynamics of these groups. Based on a careful examination of the data and analyses performed in MCK15, we propose that the divergence time estimates which they found are likely to underestimate clade ages. We believe this is due to the subset of fossil Coleoptera that MCK15 selected as calibration points, as well as the methodological approach used in their analyses. To explore the impact of fossil selection on the age of Coleoptera, we derived an alternative set of fossil calibration points based on best-practice recommendations (e.g. Parham et al., 2012),

Ultraconserved elements show utility in phylogenetic inference of Adephaga (Coleoptera) and suggest paraphyly of ‘Hydradephaga’

Abstract: The beetle suborder Adephaga has been the subject of many phylogenetic reconstructions utilizing a variety of data sources and inference methods. However, no strong consensus has yet emerged on the relationships among major adephagan lineages. Ultraconserved elements (UCEs) have proved useful for inferring difficult or unresolved phylogenies at varying timescales in vertebrates, arachnids and Hymenoptera. Recently, a UCE bait set was developed for Coleoptera using polyphagan genomes and a member of the order Strepsiptera as an outgroup. Here, we examine the utility of UCEs for reconstructing the phylogeny of adephagan families, in the first in vitro application a UCE bait set in Coleoptera. Our final dataset included 305 UCE loci for 18 representatives of all adephagan families except Aspidytidae, and two polyphagan outgroups, with a total concatenated length of 83 547 bp. We inferred trees using maximum likelihood analyses of the concatenated UCE alignment and coalescent species tree methods (astral ii, ASTRID, svdquartets). Although the coalescent species tree methods had poor resolution and weak support, concatenated analyses produced well-resolved, highly supported trees. Hydradephaga was recovered as paraphyletic, with Gyrinidae sister to Geadephaga and all other adephagans. Haliplidae was recovered as sister to Dytiscoidea, with Hygrobiidae and Amphizoidae successive sisters to Dytiscidae. Finally, Noteridae was recovered as monophyletic and sister to Meruidae. Given the success of UCE data for resolving phylogenetic relationships within Adephaga, we suggest the potential for further resolution of relationships within Adephaga using UCEs with improved taxon sampling, and by developing Adephaga-specific probes.

Phylogenomic species delimitation and host-symbiont coevolution in the fungus-farming ant genus Sericomyrmex Mayr (Hymenoptera: Formicidae): ultraconserved elements (UCEs) resolve a recent radiation

Abstract: Ants in the Neotropical genus Sericomyrmex Mayr cultivate fungi for food. Both ants and fungi are obligate, coevolved symbionts. The taxonomy of Sericomyrmex is problematic because the morphology of the worker caste is generally homogeneous across all of the species within the genus, species limits are vague, and the relationships between them are unknown. We used ultraconserved elements (UCEs) as genome-scale markers to reconstruct evolutionary history and to infer species boundaries in Sericomyrmex. We recovered an average of ∼990 UCE loci for 88 Sericomyrmex samples from across the geographical range of the genus as well as for five outgroup taxa. Using maximum likelihood and species-tree approaches, we recovered nearly identical topologies across datasets with 50–95% matrix completeness. We identify nine species-level lineages in Sericomyrmex, including two new species. This is less than the previously described 19 species, even accounting for two species for which we had no UCE samples, which brings the total number of Sericomyrmex species to 11. Divergence-dating analyses recovered 4.3 Ma as the crown-group age estimates for Sericomyrmex, indicating a recent, rapid radiation. We also sequenced mitochondrial cytochrome oxidase subunit I (COI) for 125 specimens. Resolution and support for clades in our COI phylogeny are weak, indicating that COI is not an appropriate species-delimitation tool. However, taxa within species consistently cluster together, suggesting that COI is useful as a species identification (‘DNA barcoding’) tool. We also sequenced internal transcribed spacer (ITS) and large subunit (LSU) for 32 Sericomyrmex fungal cultivars. The fungal phylogeny confirms that Sericomyrmex fungi are generalized higher-attine cultivars, interspersed with Trachymyrmex-associated fungal species, indicating cultivar sharing and horizontal transfer between these two genera. Our results indicate that UCEs offer immense potential for delimiting and resolving relationships of problematic, recently diverged species.

Phylogeny of Myrmeleontiformia based on larval morphology (Neuropterida: Neuroptera)

Abstract: The suborder Myrmeleontiformia is a derived lineage of lacewings (Insecta: Neuroptera) including the families Psychopsidae, Nemopteridae, Nymphidae, Ascalaphidae and Myrmeleontidae. In particular, Myrmeleontidae (antlions) are the most diverse neuropteran family, representing a conspicuous component of the insect fauna of xeric environments. We present the first detailed quantitative phylogenetic analysis of Myrmeleontiformia, based on 107 larval morphological and behavioural characters for 36 genera whose larvae are known (including at least one representative of all the subfamilies of the suborder). Four related families were used as outgroups to polarize character states. Phylogenetic analyses were conducted using both parsimony and Bayesian methods. The reconstructions resulting from our analyses corroborate the monophyly of Myrmeleontiformia. Within this clade, Psychopsidae are recovered as the sister family to all the remaining taxa. Nemopteridae (including both subfamilies Nemopterinae and Crocinae) are recovered as monophyletic and sister to the clade comprising Nymphidae + (Myrmeleontidae + Ascalaphidae). Nymphidae consist of two well‐supported clades corresponding to the subfamilies Nymphinae and Myiodactylinae. Our results suggest that Ascalaphidae may not be monophyletic, as they collapse into an unresolved polytomy under the Bayesian analysis. In addition, the recovered phylogenetic relationships diverge from the traditional classification scheme for ascalaphids. Myrmeleontidae are reconstructed as monophyletic, with the subfamilies Stilbopteryginae, Palparinae and Myrmeleontinae. We retrieved a strongly supported clade comprising taxa with a fossorial habit of the preimaginal instars, which represents a major antlion radiation, also including the monophyletic pit‐trap building species.

A molecular phylogeny and revised higher-level classification for the leaf-mining moth family Gracillariidae and its implications for larval host-use evolution

Abstract: Gracillariidae are one of the most diverse families of internally feeding insects, and many species are economically important. Study of this family has been hampered by lack of a robust and comprehensive phylogeny. In the present paper, we sequenced up to 22 genes in 96 gracillariid species, representing all previously recognized subfamilies and genus groups, plus 20 outgroups representing other families and superfamilies. Following objective identification and removal of two rogue taxa, two datasets were constructed: dataset 1, which included 12 loci totalling 9927 bp for 94 taxa, and dataset 2, which supplemented dataset 1 with 10 additional loci for 10 taxa, for a total of 22 loci and 16 167 bp. Maximum likelihood analyses strongly supported the monophyly of Gracillariidae and most previously recognized subfamilies and genus groups. On this basis, we propose a new classification consisting of eight subfamilies, four of which are newly recognized or resurrected: Acrocercopinae Kawahara & Ohshima subfam. n.; Gracillariinae Stainton; Lithocolletinae Stainton; Marmarinae Kawahara & Ohshima subfam. n.; Oecophyllembiinae Real & Balachowsky; Parornichinae Kawahara & Ohshima subfam. n.; Ornixolinae Kuznetzov & Baryshnikova stat. rev.; and Phyllocnistinae Zeller. The subfamily Gracillariinae is restricted to the monophyletic group comprising Gracillaria Haworth and closely related genera. We also formally transfer Acrocercops scriptulata Meyrick to Ornixolinae and use the name Diphtheroptila Vari, creating Diphtheroptila scriptulata comb. n. An exploratory mapping of larval host-use traits on the phylogeny shows strong conservation of modes of leaf mining but much higher lability of associations with host plant orders and families, suggesting that host shifts could play a significant role in gracillariid diversification. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:942814A2-DE66-41D4-8AB6-FF0B18C87EDB.

Further progress on the phylogeny of Noctuoidea (Insecta: Lepidoptera) using an expanded gene sample

Abstract: Major progress has been made recently toward resolving the phylogeny of Noctuoidea, the largest superfamily of Lepidoptera. However, numerous questions and weakly supported nodes remain. In this paper we independently check and extend the main findings of multiple recent authors by performing maximum-likelihood analyses of 5–19 genes (6.7–18.6 kb) in 74 noctuoids representing all the families and a majority of the subfamilies. Our results strongly support the six family system of Zahiri et al., with the former Lymantriidae and Arctiidae subsumed within the huge family Erebidae, and Noctuidae restricted largely to the subfamilies with so-called trifine hindwing venation. Our data also strongly corroborate monophyly of the set of four families with quadrifid forewing venation, to the exclusion of Notodontidae, and removal from the latter of Oenosandridae. Other among-family relationships, however, remain unsettled. Our evidence is equivocal on the position of Oenosandridae, which are sister group to either Notodontidae alone or to all other noctuoids. Like other recent nuclear gene studies, our results also provide no strong support for relationships among the four quadrifid forewing families. In contrast, within families our analyses significantly expand the list of robustly resolved relationships, while introducing no strong conflicts with previous molecular studies. Within Notodontidae, for which we present the largest molecular taxon sample to date, we find strong evidence for polyphyly for some, or all, recent definitions of the subfamilies Thaumetopoeinae, Pygaerinae, Notodontinae and Heterocampinae. Deeper divergences are incompletely resolved but there is strong support for multiple ‘backbone’ nodes subtending most of the subfamilies studied. Within Erebidae, we find much agreement and no strong conflict with a recent previous study regarding relationships among subfamilies, and somewhat stronger support. Although many questions remain, the two studies together firmly resolve positions for over half the subfamilies. Within Noctuidae, we find no strong conflict with previous molecular studies regarding relationships among subfamilies, but much stronger resolution along the ‘backbone’ of the phylogeny. Combining information from multiple studies yields strongly resolved positions for most of the subfamilies. Finally, our results strongly suggest that the tribes Pseudeustrotiini and Prodeniini, currently assigned to the largest subfamily, Noctuinae, do not belong there. In sum, our results provide additional corroboration for the main outlines of family-level phylogeny in Noctuoidea, and contribute toward resolving relationships within families.

Phylotranscriptomics resolves ancient divergences in the Lepidoptera

Abstract: Classic morphological studies of the oldest, so-called nonditrysian lineages of Lepidoptera yielded a well-resolved phylogeny, supported by the stepwise origin of the traits characterizing the clade Ditrysia, which contains over 98% of extant lepidopterans. Subsequent polymerase chain reaction (PCR)-based molecular studies have robustly supported many aspects of the morphological hypothesis and strongly contradicted others, while leaving some relationships unsettled. Here we bring the greatly expanded gene sampling of RNA-Seq to bear on nonditrysian phylogeny, especially those aspects that were not conclusively resolved by the combination of morphology and previous PCR-based multi-gene studies. We analysed up to 2212 genes in each of 28 species representing all 12 superfamilies and 15 of 21 families of nonditrysians, plus trichopteran outgroups and representative Ditrysia. Our maximum likelihood phylogeny estimates used both nonsynonymous changes only (degen1 coding) and all nucleotides (nt123) partitioned by codon position, recovering a novel hypothesis for early glossatan relationships that is the most strongly supported to date. We find strong support for Micropterigidae alone as the sister group to all other Lepidoptera, in agreement with morphology and early molecular evidence, but in contrast to recent PCR-based studies. Also very strongly supported are the previously recognized clades Angiospermivora, Heteroneura, Eulepidoptera and Euheteroneura. Finally, we find strong support for paraphyly of the southern hemisphere family Palaephatidae, with the South American genus Palaephatus Butler forming the previously undetermined sister group to Ditrysia. The remaining palaephatids, Australian and South American, form the sister group to Tischeriidae.

Phylogeny and reclassification of Cryptini (Hymenoptera, Ichneumonidae, Cryptinae), with implications for ichneumonid higher-level classification

Abstract: The first comprehensive phylogenetic study of the wasp tribe Cryptini (Hymenoptera, Ichneumonidae, Cryptinae) is presented, based on 109 morphological characters and molecular data from seven loci The dataset includes 370 species, 308 of which are from Cryptini, covering 182 of its 250 genera Results from parsimony and likelihood analyses are generally congruent The topology has several implications for ichneumonid higher-level classification Previous definitions of the Ichneumoniformes clade are supported, though newly including the Microleptinae The cryptine subtribe Ateleutina is consistently recovered outside of the Cryptini clade and should be treated as a separate subfamily, Ateleutinae statn The tribe Phygadeuontini is shown to be polyphyletic: while most of the sampled taxa were recovered in a single clade, many of its members are more closely related to the Ichneumoninae, Ateleutinae or Cryptini Pending a more detailed study, the group should be treated as a separate subfamily, Phygadeuontinae stat rev The former Hemigastrini are recovered as largely monophyletic but with important exceptions Hemigaster Brulle is recovered as part of the Phygadeuontini and is transferred to that group Echthrus Gravenhorst is consistently recovered as part of Cryptini, rendering Aptesini as the correct name for the tribe The subfamily Cryptinae should be restricted to the tribes Aptesini and Cryptini Within Cryptini, the results show little support for the current subtribal classification, with most subtribes recovered as polyphyletic A number of relatively stable clades are identified and discussed, but the relationships among them are weakly supported Most of these clades are morphologically heterogeneous and building a subtribal classification based on them would be ineffectual; they are therefore treated under the informal designation of genus groups The results highlight the ubiquity of morphological homoplasy in Cryptini, and provide a framework from which to address further systematic and evolutionary questions on this hyperdiverse group of parasitic wasps

A new genus of hell ants from the Cretaceous (Hymenoptera: Formicidae: Haidomyrmecini) with a novel head structure

Abstract: An unusual Cretaceous trap jaw ant is described from Burmese amber dated to the Late Cretaceous. Linguamyrmex vladi gen.n. sp.n. is distinguished by an unusual suite of morphological characters indicating specialized predatory behaviour and an adaptive strategy no longer found among modern ant lineages. The clypeus, highly modified as in other closely related haidomyrmecine hell ants, is equipped with a paddle-like projection similar to Ceratomyrmex. X-ray imaging reveals that this clypeal paddle is reinforced, most probably with sequestered metals. Presumably this fortified clypeal structure was utilized in tandem with scythe-like mandibles to pin and potentially puncture soft-bodied prey. This unique taxon, which stresses the diversity of stem-group ants, is discussed in the context of modern and other Cretaceous trap jaw ant species. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:40D636A3-4D88-470A-BC5B-85ABFD1A49E2.

Citation of taxonomic publications: the why, when, what and what not

Abstract: Species-level research in taxonomy is inadequately cited (Krell, 2000; Agnarsson & Kuntner, 2007; Ebach et al., 2011; Haszprunar, 2011; Wägele et al., 2011). This has been recognized for many decades, but it did not hurt taxonomy until citation counts started to influence funding, careers, and positions (Krell, 2000; Agnarsson & Kuntner, 2007; Bortolus, 2008; Seifert et al., 2008). One seemingly easy fix that has been proposed or even mandated by some journals is to require full citations of original species descriptions (see also Seifert et al., 2008; Haszprunar, 2011). For example, Organisms, Diversity, and Evolution ‘will now require that author citations for species names be supported by inclusion of the full, accompanying reference’ (Bininda-Emonds, 2011) and Zootaxa’s instructions for authors includes: ‘Author(s) of species name must be provided when the scientific name of any animal species is first mentioned (the year of publication needs not be given; if you give it, then provide a full reference of this in the reference list)’. Other journals encourage or allow such citations (see Wägele et al., 2011). Here, I argue that citing original descriptions is often misleading and that the proper citation credits should instead go to the literature that was used for specimen identification and the publications that contain the most accurate information on the currently accepted species limits (‘taxon concept’ sensu Franz, 2005) of the species under study (see also Agnarsson & Kuntner, 2007). I argue that biological publications need to include identification techniques and taxon concepts in the ‘Methods’ section and that the identifications should be treated as a ‘Result’ in the corresponding section. The information should be succinctly presented so that reviewers and editors can assess compliance.

Early lineages of Vespidae (Hymenoptera) in Cretaceous amber

Abstract: Three wasp (Hymenoptera: Vespidae) fossils in Cretaceous amber (Late Albian) of northern Myanmar are described. Two are new species of the Mesozoic genus Curiosivespa (Rasnitsyn): C. zigrasi sp.n. and C. striata sp.n. The third species, Protovespa haxairei gen.n. et sp.n., has a combination of features unique among Mesozoic Priorvespinae and the extant subfamilies. These well preserved fossils provide new morphological data for a cladistic analysis of the basal lineages of Vespidae. Results suggest that Euparagiinae is the sister group of all other Vespidae. The new genus Protovespa appears more closely related to extant Masarinae, Eumeninae and social wasps than to Priorvespinae. We assign it to a new subfamily: Protovespinae. Finally, fossil information combined with a phylogenetic tree shows that the main groups of Vespidae probably evolved during the Early Cretaceous. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:1E7E4796-6E70-4D81-BB34-0FEEA765DC25.

A revision of the Encarsia pergandiella species complex (Hymenoptera: Aphelinidae) shows cryptic diversity in parasitoids of whitefly pests

Abstract: Encarsia pergandiella Howard, described from North America (USA), and Encarsia tabacivora Viggiani, described from South America (Brazil) (Hymenoptera: Aphelinidae), are two formally recognized taxonomic entities, that have been treated by several authors as synonyms due to lack of strong diagnostic characters. Taxonomy of these species is further complicated because several populations, geographically separated and differing in their biology, have been included under the concept of E. pergandiella. Among these, a population originally collected in Brazil and introduced to North America reproduces by thelytokous parthenogenesis and is infected by the symbiont Cardinium, while a morphologically indistinguishable population, naturally occurring in Texas, is biparental and infected by a related strain of Cardinium that induces cytoplasmic incompatibility. A third population known from California and introduced to the Old World is biparental and uninfected by intracellular symbionts. While adult females of the first two populations have entirely light yellow bodies and pupate face up (light form), those of the third population have largely brown bodies and pupate face down (dark form). Other dark form populations are known from Texas, Florida and New York. Because these parasitoids are economically important biological control agents of cosmopolitan whitefly pests, it is critical to characterize them correctly. In this study, we integrated molecular and morphometric analyses to substantiate observed differences in biological traits, and resolve the complicated taxonomy of this species complex. We sequenced the mitochondrial cytochrome c oxidase subunit I gene and the D2 region of the ribosomal 28S gene for individuals of both light form (from Texas and Brazil) and dark form (from California, Texas, Italy and Canary Islands) originating from laboratory cultures or collected in the field. Phylogenetic analysis unambiguously distinguished three well-supported groups corresponding to the Texas light form, the Brazil light form and the dark form. Individuals of these three groups, in combination with all available type material (E. pergandiella, its synonym Encarsia versicolor Girault and E. tabacivora) and additional museum specimens of the dark form from New York and Italy, were subjected to multivariate morphometric analyses using Burnaby principal component analysis followed by a linear discriminant analysis, and multivariate ratio analysis. Overall, the analyses showed that: (i) E. pergandiella and E. tabacivora are two distinct species; (ii) the thelytokous Brazil light form corresponds to E. tabacivora; (iii) the biparental Texas light form is a new species formally described here as Encarsia suzannae sp.n.; (iv) two new biparental species can be referred to the dark form, one described as Encarsia gennaroi sp.n. including the populations sampled in California, Texas, Italy and Canary Islands, and the other corresponding to the population from New York described as Encarsia marthae sp.n. A dichotomous key for both sexes of the species of the E. pergandiella complex is provided for identification. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:278475A0-C2C4-4400-A042-A5716457829D.

Higher-level phylogeny of diving beetles (Coleoptera: Dytiscidae) based on larval characters

Abstract: A comprehensive higher-level phylogeny of diving beetles (Dytiscidae) based on larval characters is presented. Larval morphology and chaetotaxy of a broad range of genera and species was studied, covering all currently recognized subfamilies and tribes except for the small and geographically restricted Hydrodytinae, where the larva is unknown. The results suggest several significant conclusions with respect to the systematics of Dytiscidae including the following: monophyly of all currently recognized subfamilies, although Dytiscinae when considered in a broad context is rendered paraphyletic by Cybistrinae; currently recognized tribes are monophyletic except for Agabini, Hydroporini and Laccornellini; inter-subfamily and inter-tribe relationships generally show weak support, except for a few well supported clades; three distinct clades are recognized within Dytiscinae [Dytiscini sensu lato (i.e. including the genera Dytiscus Linnaeus and Hyderodes Hope), Hydaticini sensu lato, and Cybistrini]; and recognition of Pachydrini as a distinct tribe. Other less robust results include: Methlini sister to the rest of Hydroporinae; relative basal position of Laccornini, Hydrovatini and Laccornellini within Hydroporinae; close relationship of Agabinae and Copelatinae; Matinae nested deep within Dytiscidae, as sister to a large clade including Colymbetinae, Coptotominae, Lancetinae and Dytiscinae sensu lato; the sister-group relationship of Agabetini and Laccophilini is confirmed. The results presented here are discussed and compared with previous phylogenetic hypotheses based on different datasets, and the evolution of some significant morphological features is discussed in light of the proposed phylogeny. All suprageneric taxa are diagnosed, including illustrations of all relevant synapomorphies, and a key to separate subfamilies and tribes is presented, both in traditional (paper) format and as an online Lucid interactive identification key.

Taxonomy as a hypothesis: testing the status of the Bermuda buckeye butterfly Junonia coenia bergi (Lepidoptera: Nymphalidae)

Abstract: Species determination and definition in eukaryotes have traditionally been based on morphology, with little focus on genetic differentiation Molecular methods allow for the independent assessment of morphology-based taxonomic hypotheses Three criteria used to define a full species for taxonomic purposes are morphological distinction, formation of a monophyletic lineage, and reproductive isolation Junonia butterflies (Nymphalidae) are becoming an important experimental model system, but the taxonomy of many New World Junonia species is unclear One of these species is J coenia, which contains the subspecies J coenia coenia, J coenia grisea and J coenia bergi Previous studies suggest that J coenia grisea may meet the criteria for full species status Therefore, we evaluated the geographically isolated and rarely studied Bermuda buckeye butterfly J coenia bergi to determine if it was similarly distinct Physical examination of specimens and phylogenetic and population genetic analyses of mitochondrial cytochrome c oxidase subunit I, nuclear wingless, and complete mitochondrial genome sequences suggest that while J coenia bergi is smaller in body size than many Junonia and has distinctive ventral hindwing colouration, it does not form a monophyletic lineage and shows indications of continued gene flow with North American mainland J coenia coenia populations Thus, J coenia bergi does not meet the criteria for full species designation, but geographic isolation, morphological distinctiveness, and cultural importance suggest that it remain recognized as a subspecies of J coenia Similar analyses will be useful for addressing further taxonomic questions in Junonia and other taxa, especially where morphology-based taxonomic determinations are ambiguous

The phylogeny of lance lacewings (Neuroptera: Osmylidae)

Abstract: The first phylogeny of the lacewing family Osmylidae is presented here based on a total evidence analysis of DNA sequences for multiple gene loci and morphology for representatives of almost all extant genera. Our phylogeny shows a basal dichotomy in the family, with subfamilies Protosmylinae, Spilosmylinae and Gumillinae comprising one lineage, and the other lineage including Osmylinae, Porisminae, Eidoporisminae, Kempyninae and Stenosmylinae. The status of Paryphosmylus Kruger and Lysmus Navas as members of Protosmylinae is affirmed as well as the placement of Gumillinae near Protosmylinae and Spilosmylinae. Our results suggest that Porisminae, Eidoporisminae and Stenosmylinae evolved from a common ancestor, and their relationships, including likely paraphyly of Stenosmylinae, requires further assessment. Divergence time analysis revealed that the family originated during the Late Permian before the break-up of the supercontinent Pangaea and that present generic distributions are not due to Gondwanan biogeographic events. All major subfamily-level lineages were present by the end of the Triassic, in agreement with the rich Mesozoic-aged fossil record for the family.

Phylogeny, classification and divergence times of pygmy leaf‐mining moths (Lepidoptera: Nepticulidae): the earliest lepidopteran radiation on Angiosperms?

Abstract: Nepticulidae represent one of the early diverging Lepidoptera lineages, and the family currently comprises over 850 described species The larvae of the vast majority of the species are leaf miners on Angiosperms and highly monophagous, which has led to persistent ideas on coevolution with their plant hosts We present here a molecular phylogeny based on eight gene fragments from 355 species, representing 20 out of 22 extant Nepticulidae genera Using two fossil calibration points, we performed molecular dating to place the origin of the family in the Early Cretaceous, before the main Angiosperm diversification Based on our results we propose a new classification, abandoning all ranks between family and genus, as well as subgenera to allow for a stable classification The position of Enteucha Meyrick within Nepticulidae remains somewhat ambiguous, and the species-rich cosmopolitan genus Stigmella Schrank, with nearly half of all described Nepticulidae, requires further study Ectoedemia Busck, Zimmermannia Hering, Acalyptris Meyrick, Etainia Beirne, Parafomoria Borkowski, Muhabbetana Kocak & Kemal and Fomoria Beirne appear to have diversified in a relatively short evolutionary period, leading to short branches in the molecular phylogeny and unclear suprageneric relations Otherwise support values throughout the phylogeny are mostly high and the species groups, genera and higher clades are discussed in respect of their supporting morphological and life-history characters Wing venation characters are confirmed to be mostly reliable and relevant for Nepticulidae classification, but some other previously used characters require reinterpretation The species groups of most genera are recovered, but only partly so in the large genus Stigmella The molecular dating results are compared with existing knowledge on the timing of the Angiosperm radiation and reveal that the diversification of Nepticulidae could largely have been contemporaneous with their hosts, although some of the genera restricted to a single plant family appear to have begun to diversify before their hosts

Interrelationships and diversification of Argynnis Fabricius and Speyeria Scudder butterflies

Abstract: Diverse radiations of insects are often associated with adaptations to host plants, and well-resolved phylogenetic relationships are required to fully understand them. Palearctic Argynnis and related subgenera, together with North American Speyeria butterflies make up a radiation whose species hypotheses are confounded by shared wing colour patterns between sympatric populations of closely related recognized species. Previous studies of this group indicate that Speyeria is a lineage within Argynnis, but sampling in these studies has either involved too few Speyeria species or incomplete sampling of Argynnis species. Thus, no comprehensive phylogenetic analysis exists for all members that answers the question of monophyly of Speyeria, or other subgeneric taxa, and their relationship to Argynnis species. We completed a phylogenetic analysis of all North American Speyeria species and all but one species within Argynnis, using one mitochondrial (cytochrome c oxidase I, COI) and four nuclear genes [elongation factor 1 alpha (EF1α), wingless (WG), glyceraldehyde-3-phosphate dehydrogenase (GAPDH), and ribosomal protein S5 (RPS5)]. The results indicate three major lineages within Argynnis s.l.: two Palearctic and one containing both Palearctic and Nearctic species. In summary, the phylogenetic analyses suggest the need for reorganization into three natural groups: Argynnis, Fabriciana and Speyeria. Within each of these genera the phylogenetic hypothesis indicates an evolutionary history marked by rapid diversification and potential extinction, followed by ongoing lineage sorting. The position of North American Speyeria is nested within the Palearctic lineages, which indicates that the radiation began in Asia and was fuelled by existing Viola diversity in North America. Dating analyses of Viola and Speyeria corroborate this hypothesis. The current North American Speyeria species are mixed on the tree, indicating a recent and ongoing radiation. These results provide needed clarity on the evolution of this group, which contains species of conservation concern.

Two's company, three's a crowd: new insights on spruce budworm species boundaries using genotyping-by-sequencing in an integrative species assessment (Lepidoptera: Tortricidae)

Abstract: Species delimitation requires an assessment of varied traits that can contribute to reproductive isolation, as well as of the permanence of evolutionary differentiation among closely related lineages. Integrative taxonomy, including the combination of genome-wide molecular data with ecological data, offers an effective approach to this issue. We use genotyping-by-sequencing together with a review of ecological divergence to assess the traditionally recognized species status of three closely related members of the spruce budworm species complex, Choristoneura fumiferana (Clemens), C. occidentalis Freeman (=C. freemani Razowski) and C. biennis Freeman, each of which is a major defoliator of conifer forests. We sampled a broad region of overlap between these three taxa in Alberta and British Columbia (Canada) where potential for gene flow provides a strong test of the durability of divergence among lineages. A total of 2218 single nucleotide polymorphisms (SNPs) were assayed, and patterns of differentiation were evaluated under the biological, ecological, genotypic cluster and phylogenetic species concepts. Choristoneura fumiferana was genetically distinct with substantial barriers to genetic exchange with C. occidentalis and C. biennis. Conversely, divergence between C. occidentalis and C. biennis was limited to a small subset of outlier loci and was within the range observed within any one of the taxa. Considering both population genetic and ecological patterns of divergence, C. fumiferana should continue to be recognized as a distinct species, and C. biennis (syn.n.) should be treated as a subspecies (C. occidentalis biennis Freeman, 1967) of C. occidentalis, thereby automatically establishing the nominate name C. occidentalis occidentalis Freeman, 1967 for univoltine populations of this species.

Integrative taxonomy versus taxonomic authority without peer review: the case of the Oriental fruit fly, Bactrocera dorsalis (Tephritidae)

Abstract: 1. Major global horticultural and food security tephritid fruit fly pests, Bactrocera papayae (papaya fruit fly) and B. invadens (invasive fruit fly), were synonymised with B. dorsalis (Oriental fruit fly) by Schutze et al. (2015a) based on extensive integrative taxonomic evidence from multiple sources. This synonymy was peer reviewed by eight independent experts. 2. Drew & Romig (2016) withdrew B. papayae and B. invadens from synonymy based on opinion drawn primarily from disparate geographical distribution, morphological, and host use information. This reversal was not subjected to peer review. 3. We consider the withdrawal from synonymy as invalid due to significant errors and misrepresentations of the literature provided in the arguments of Drew & Romig (2016) that we propose would not have withstood peer scrutiny. 4. This case reflects a broader issue of individual taxonomic authorities using opinion to challenge extensive evidence generated via scientific hypothesis-testing methods by discipline specialists. 5. We recommend that taxonomic acts not subjected to peer review, especially of pest species, be actively discouraged by the broader scientific and regulatory community.

Phylogeny of pleasing lacewings (Neuroptera: Dilaridae) with a revised generic classification and description of a new subfamily

Abstract: The phylogeny of pleasing lacewings (Neuroptera: Dilaridae) is reconstructed for the first time based on morphological data using all fossil and extant genera. Accordingly, a revised generic classification of Dilaridae is proposed, with a new subfamily (i.e. Berothellinae subfam.n.) erected based on its remarkably different morphological features from the other dilarid subfamilies. A revision of all dilarid genera is presented, including descriptions of some little-known species from Asia and Mid-Cretaceous Burmese amber. New genera and species herein described include Berothella holzschuhi U. Aspock, Liu & H. Aspock, sp.n., Cretodilar burmanus Liu & Zhang, gen. et sp.n., Dilar cretaceus Liu & Zhang, sp.n., Neonallachius orientalis Liu, U. Aspock & H. Aspock, sp.n. and Neonallachius thailandicus Liu & Winterton, sp.n. Two new combinations, i.e. Neonallachius krooni (Minter, 1986), comb.n. and Neonallachius ponomarenkoi (Zakharenko, 1991), comb.n., are proposed. Evolutionary patterns of some important characters and the historical biogeography of Dilaridae are also discussed. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:68836312-FBDC-4F9F-8516-2365F44596BF.

Phylogeography and taxonomy of the Psorodonotus caucasicus (Orthoptera, Tettigoniidae) group: independent double invasion of the Balkans from the Caucasus

Abstract: High mountain ranges within and around Anatolia have played an important role in the biogeography of mountainous organisms, which are threatened by climate change. Using the Psorodonotus caucasicus species group (PCG), whose range is the Caucasus, Anatolia and the Balkans, as a model group, this study aims to determine the importance of Anatolian mountain ranges and to estimate possible future effects of global warming on such lineages by establishing an analogy between interglacial dynamics and global warming. Twenty populations, representing all extant species of the PCG, were studied. Sequences of four mitochondrial loci and three nuclear loci were obtained and used in phylogenetic, time estimation and phylogeographic analyses of PCG members. Additionally, ecological modelling analyses were conducted to estimate the range of PCG members at present and during the last glacial maximum and the last interglacial, with the aim of determining the roles of altitudes and glacial cycles in shaping the distribution of the PCG. Phylogenetic analyses supported the monophyly of the PCG as a whole and each individual species with the exception of P. suphani. The radiation of the PCG started approximately 3 Ma, according to beast. Species distribution models suggested a wider range during the last glacial maximum than during the last interglacial and at present, but these models also suggested that present conditions represent a substantial range loss. The PCG dispersed to the Balkans through two independent corridors from the Caucasus. The following conclusions were reached based on the results: (i) the uplift of the Pontic and East Anatolian Blocks during the mid‐Pliocene and the closing of the Mediterranean and the Paratethys connection in the Plio–Pleistocene and the mid‐Pleistocene Transition are the main events that triggered the radiation of the PCG; (ii) the PCG radiated from a Caucasian origin and invaded the Balkans twice, first through Northern Anatolia and second via the ‘Taurus Way’ dispersal corridor; (iii) lowlands are a barrier to the dispersal of PCG members; (iv) the last four intense glacial periods had little effect on the radiation of the PCG; (v) the range of the PCG has receded substantially from its original southern margin, and P. salmani and P. anatolicus have become extinct since the 1950s, possibly because of global warming; and (vi) automatic species delimitation tests supported species status of the morphospecies P. caucasicus, P. macedonicus, P. fieberi, P. illyricus and P. ebneri, but not that of P. suphani, thus, P. suphani Taylan & Sirin, 2014 syn.n. synonymized with P. caucasicus (Fischer von Waldheim, 1846).

Integrated species delimitation and conservation implications of an endangered weevil Pachyrhynchus sonani (Coleoptera: Curculionidae) in Green and Orchid Islands of Taiwan

Abstract: Oceanic islands are productive habitats for generating new species and high endemism, which is primarily due to their geographical isolation, smaller population sizes and local adaptation. However, the short divergence times and subtle morphological or ecological divergence of insular organisms may obscure species identity, so the cryptic endemism on islands may be underestimated. The endangered weevil Pachyrhynchus sonani Kono (Coleoptera: Curculionidae: Entiminae: Pachyrhynchini) is endemic to Green Island and Orchid Island of the Taiwan-Luzon Archipelago and displays widespread variation in coloration and host range, thus raising questions regarding its species boundaries and degree of cryptic diversity. We tested the species boundaries of P. sonani using an integrated approach that combined morphological (body size and shape, genital shape, coloration and cuticular scale), genetic (four genes and restriction site-associated DNA sequencing, RAD-seq) and ecological (host range and distribution) diversity. The results indicated that all the morphological datasets for male P. sonani, except for the colour spectrum, reveal overlapping but statistically significant differences between islands. In contrast, the morphology of the female P. sonani showed minimum divergence between island populations. The populations of P. sonani on the two islands were significantly different in their host ranges, and the genetic clustering and phylogenies of P. sonani established two valid evolutionary species. Integrated species delimitation combining morphological, molecular and ecological characters supported two distinct species of P. sonani from Green Island and Orchid Island. The Green Island population was described as P. jitanasaius sp.n. Chen & Lin, and it is recommended that its threatened conservation status be recognized. Our findings suggest that the inter-island speciation of endemic organisms inhabiting both islands may be more common than previously thought, and they highlight the possibility that the cryptic diversity of small oceanic islands may still be largely underestimated.

Multilocus species delimitation in Mesoamerican Scaptotrigona stingless bees (Apidae: Meliponini) supports the existence of cryptic species

Abstract: Accelerating taxonomic knowledge and making accurate species identifications are critically important given the current biodiversity crisis, particularly in biodiversity hotspots such as Mesoamerica. Objective species delimitation that reduces investigator‐driven bias is fundamental to the establishment of appropriate conservation strategies, above all in managed species. Previous morphological and molecular studies on three managed stingless bee species of the genus Scaptotrigona distributed in Mexico (S. mexicana, S. pectoralis and S. hellwegeri) suggested that both S. mexicana and S. hellwegeri are cryptic species complexes. Herein we tested species delimitation by analysing sequence information of five markers (two mitochondrial: cox1 and 16S, and three nuclear: ITS1, EF1‐α, ArgK) within a Bayesian coalescent framework to test the putative species. We obtained two different hypotheses using a Generalized Mixed Yule Coalescent (GMYC) model: four (cox1) and six (16S) species. After the species validation step with the Bayesian species‐delimitation analysis (BPP), we suggest that only S. mexicana is a complex of two species with different distribution (along the Pacific and the Atlantic coasts, respectively). We highly recommend avoiding colony exchange between geographical regions in order to conserve the genetic integrity of both taxa.

A phylogeny of the treehopper subfamily Heteronotinae reveals convergent pronotal traits (Hemiptera: Auchenorrhyncha: Membracidae)

Abstract: Even within an insect family famous for its morphological diversity, the treehopper subfamily Heteronotinae is a microcosm of pronotal variation, displaying remarkably dissimilar thoracic ornamentations among its ten included genera. Presented here is a reconstruction of heteronotine relationships based on DNA nucleotide sequence data from five nuclear and two mitochondrial genes from a comprehensive sample of ingroup taxa (including exemplars of all genera except for the monotypic Dysyncritus Fowler, 1895). Concordant phylogenetic estimates support the monophyly of Heteronotinae sensu stricto (i.e. excluding Darnoides Fairmaire, 1846), as well as the monophyly of the included genera Heteronotus Laporte, 1832, Nassunia Stål, 1862, Omolon Walker, 1862, Rhexia Stål, 1867, and Smiliorachis Fairmaire, 1846. Conversely, Anchistrotus Buckton, 1902, Allodrilus Evangelista, 2014 and Iria Stål, 1867 were not recovered as monophyletic, although topology comparison tests failed to reject the nonmonophyly of the latter two genera, nor Heteronotinae, as currently circumscribed. These results are interpreted in the context of available morphological evidence, which generally supports these findings. Also addressed here is the evolution of selected pronotal features representing major aspects of heteronotine morphology. Bayesian stochastic mapping of pronotal traits estimated multiple character state transitions, suggesting repeated acquisition of pronotal features, most of which have evolved independently within the diverse genus Heteronotus. These convergences may provide insight into the role of the pronotum in heteronotine groups that appear to serve different adaptive strategies, such as mimicry, crypsis and defence against predators. Furthermore, our results show a substantial diversity in need of formal description, including possible new genera, and several unique sexually dimorphic syndromes that may constitute a species complex within Heteronotus delineatus Walker, 1858. Nomenclatural changes are proposed for Nassunia nigromacula (Funkhouser, 1940) new combination and Smiliorachis inornata Stål, 1862 combination reinstated, which were found to be misplaced at the generic and subfamily levels according to our analyses.

Molecular phylogeny of Sericostomatoidea (Trichoptera) with the establishment of three new families

Abstract: We inferred the phylogenetic relationships among 58 genera of Sericostomatoidea, representing all previously accepted families as well as genera that were not placed in established families. The analyses were based on five fragments of the protein coding genes carbamoylphosphate synthetase (CPSase of CAD), isocitrate dehydrogenase (IDH), Elongation factor 1a (EF-1a), RNA polymerase II (POL II) and cytochrome oxidase I (COI). The data set was analysed using Bayesian methods with a mixed model, raxml, and parsimony. The various methods generated slightly different results regarding relationships among families, but the shared results comprise support for: (i) a monophyletic Sericostomatoidea; (ii) a paraphyletic Parasericostoma due to inclusion of Myotrichia murina, leading to synonymization of Myotrichia with Parasericostoma; (iii) a polyphyletic Sericostomatidae, which is divided into two families, Sericostomatidae sensu stricto and Parasericostomatidae fam.n.; (iv) a polyphyletic Helicophidae which is divided into Helicophidae sensu stricto and Heloccabucidae fam.n.; (v) hypothesized phylogenetic placement of the former incerta sedis genera Ngoya, Seselpsyche and Karomana; (vi) a paraphyletic Costora (Conoesucidae) that should be divided into several genera after more careful examination of morphological data; (vii) reinstatement of Gyrocarisa as a valid genus within Petrothrincidae. A third family, Ceylanopsychidae fam.n., is established based on morphological characters alone. A hypothesis of the relationship among 14 of the 15 families in the superfamily is presented. A key to the families is presented based on adults (males). Taxonomic history, diagnosis, habitat preference and distribution data for all sericostomatoid families are presented. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:CF6A6B9F-6A72-4265-BD09-3A710DFCD7B1.

Defence by plant toxins in milkweed bugs (Heteroptera: Lygaeinae) through the evolution of a sophisticated storage compartment

Abstract: Species of the heteropteran subfamily Lygaeinae possess special subcuticular compartments to store cardiac glycosides, plant-derived defensive compounds, which they release upon predator attack. In adults of the large milkweed bug, Oncopeltus fasciatus, these storage compartments have previously been described as a modified integument, forming a fluid-filled dorsolateral space. Here we use three-dimensional imaging of serial histological sections and synchrotron radiation-based micro-computed tomography data to reveal the morphology of these storage compartments and the mechanisms used for the release of a cardiac glycoside-rich fluid upon attack. Our comparative analysis revealed that the morphology and release mechanism vary among the species investigated. By reconstructing these traits on a recent molecular phylogeny of the Lygaeinae, we demonstrate that the adaptations for the storage and release of cardiac glycosides have evolved in a stepwise manner.

Biogeographic mirages? Molecular evidence for dispersal-driven evolution in Hydrobiusini water scavenger beetles

Abstract: Water beetles of the tribe Hydrobiusini are globally distributed in the northern hemisphere and all austral continents except Antarctica. A remarkable clade also occurs in the Hawaiian Islands. The phylogenetic relationships among genera were recently investigated using a combination of molecules and morphology. Here, we use this phylogenetic framework to address the biogeographic evolution of this group using Bayesian fossil-based divergence times, and model-based maximum likelihood ancestral range estimations. We recover an origin of the tribe in the Cretaceous ca. 100 Ma. Our biogeographic analyses support an origin of the tribe in Laurasia followed by the colonization of Australia. However, a Gondwanan origin of the group cannot be ruled out when considering the fossil record. The timeframe of the tribe's evolution as well as the model-based approach of ancestral range estimation favour a scenario invoking multiple transoceanic dispersal events over a Gondwana vicariance hypothesis. The Hawaiian radiation originated from long-distance dispersal to now-submerged islands, paired with dispersal to new islands as they formed.

To be or not to be a valid genus: the systematic position of Ophyra R.‐D. revised (Diptera: Muscidae)

Abstract: Ophyra Robineau-Desvoidy is one of the better-studied genera of the family Muscidae (Diptera) The larvae of species of this genus feed on highly decomposed organic matter of various origins, and may reveal predatory behaviour as they mature These feeding habits, combined with the widespread distribution and close association with human dwellings of some species, give the genus commercial and medico-legal importance However, the systematic position of Ophyra has been a matter of debate for many years Ophyra has been considered by muscid workers to be either a valid genus or a junior synonym of Hydrotaea Robineau-Desvoidy A lack of agreement about the systematic position of Ophyra has led to serious errors, particularly in the applied literature Recent molecular and morphological studies provided contradictory information on the validity of the genus and its subfamilial classification We revise the systematic position of Ophyra herein by means of molecular phylogenetic reconstruction Our results are incongruent with opinions on the systematic position of Ophyra based on previously published molecular phylogenies, although they correspond with the concept of the genus based on adult morphology All analyses of the concatenated dataset revealed Ophyra as monophyletic and placed within a paraphyletic Azeliini Depending on the phylogenetic approach, Ophyra was placed within either a monophyletic, paraphyletic or polyphyletic Hydrotaea, yet always closely related to a certain group of species We conclude that Ophyra, as currently defined, should not be considered a valid name, yet a detailed reconstruction of the genus Hydrotaea awaits future studies with greater increases in both taxon sampling and the number of molecular markers

‘Back to Africa’: increased taxon sampling confirms a problematic Australia‐to‐Africa bee dispersal event in the Eocene

Abstract: Colletidae is a predominantly southern hemisphere bee family with a Late Cretaceous origin and with an inferred ancestral region covering late Gondwanan South America, Antarctica and Australia. One highly diverse colletid subfamily, Euryglossinae, is entirely restricted to Australia and the strictly Afrotropical subfamily Scrapterinae has been inferred as its sister clade. This has led to suggestions that Scrapterinae represents a highly unusual post-Gondwanan dispersal from Australia to Africa, but phylogenetic studies to date have included only minimal representatives from each subfamily. Here we greatly increase the level of species sampling of both subfamilies and develop a molecular phylogeny based on one mitochondrial and two nuclear genes. Our results indicate that the broad results of earlier studies are robust to substantially greater taxon sampling, and we infer a divergence date between the two subfamilies in the early Eocene. Dispersal pathways between Africa and Australia during that time are problematic, with several studies suggesting dispersals via the now largely submerged Kerguelen and Crozet Plateaus. Our results contribute another example of a puzzling sister-clade relationship between African and Australian taxa and indicate the need to better understand southern hemisphere subaerial configurations, including Antarctica, and ocean and wind currents at those times.

Phylogeny of the tribe Naupactini (Coleoptera: Curculionidae) based on morphological characters.

Abstract: Naupactini (Curculionidae: Entiminae) is a primarily Neotropical tribe of broad-nosed weevils with its highest genus and species diversity in South America. Despite several taxonomic contributions published during the last decades, the evolutionary history of Naupactini remains poorly understood. We present the first comprehensive phylogenetic analysis for this tribe based on a data matrix of 100 adult morphological characters scored for 70 species, representing 55 genera of Naupactini (ingroup) and four outgroups belonging to the entimine tribes Otiorhynchini, Entimini, Eustylini and Tanymecini. According to the most parsimonious tree Artipus does not belong to Naupactini; the genera with flat and broad antennae, formerly assigned to other entimine tribes, form a monophyletic group (Saurops (Curiades (Aptolemus (Platyomus)))) related to the clade (Megalostylus (Megalostylodes (Chamaelops Wagneriella))); and the genera distributed along the high Andes, Paramos and Puna form a natural group (Asymmathetes (Amphideritus (Leschenius (Amitrus (Obrieniolus (Melanocyphus Trichocyphus)))))), nested within a larger clade that includes Pantomorus, Naupactus and allied genera. Atrichonotus, Hoplopactus, Mimographus and Naupactus are not recovered as monophyletic. In order to address the taxonomic implications of our phylogenetic analysis, we propose the following nomenclatural changes: to transfer Artipus from Naupactini to Geonemini, to revalidate the genera Mimographopsis (type species M. viridicans), and to revalidate the genus Floresianus (type species F. sordidus). The evolution of selected characters is discussed. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:C8AA4388-A2F0-4E2D-889A-500BEA5A9DE1.